Thomas F. Schilling

Professor and Chair, Developmental & Cell Biology
School of Biological Sciences

Ph.D., University of Oregon, Eugene, Oregon, 1993, Developmental Biology

M.S., University of Michigan, Ann Arbor, Michigan, 1987, Neurobiology

B.S., Davidson College, Davidson, North Carolina, 1985, Biology

Phone: (949) 824-2479
Fax: (949) 824-4709

University of California, Irvine
4109 Natural Sciences II
4462 Natural Sciences II
Mail Code: 2300
Irvine, CA 92697

picture of Thomas F. Schilling

zebrafish, development, neural crest, segmentation, hindbrain, retinoic acid, endothelin, pattern formation, embryogenesis, morphogenesis
URLs UCI Faculty Web Site
UCI Lab Web Site
Pew Scholar in Biomedical Sciences
Fellow of the American Association for the Advancement of Science (AAAS)
Appointments University College London and Imperial Cancer Research Fund, London, England
Our laboratory uses genetics and molecular biology to study pattern formation in the zebrafish embryo. The rapid development and simple anatomy of this teleost embryo, together with recently developed techniques for reverse genetics and a nearly complete genome sequence, make zebrafish a powerful molecular genetic system for studying the mechanisms of development. We are interested in how gene functions translate into cell behaviours and the formation of tissues and organs. We focus on two main areas:
1) Neural crest specification and formation of the craniofacial skeleton,
2) Cell interactions and formation of the anterior-posterior axis of the nervous system.
Publications Wang W, Holmes WR, Sosnik J, Schilling TF and Nie Q (2017). Cell sorting and noise-induced plasticity coordinate to sharpen boundaries between gene expression domains. PLoS Computational Biology 13(1):e1005307. doi: 10.1371/journal.pcbi.1005307.
  Muto A and Schilling TF (2017). Zebrafish as a model to study cohesin and cohesinopathies.
In K Yokomori and Katsuhiko Shirahige (Eds.). Cohesin and Condensin: Methods and Protocols. Methods in Mol Biol. Vol 1515, pp. 177-196 Springer Press.
  Vibert L, Aquino G, Gehring I, Subkhankulova T, Schilling TF, Rocco A and Kelsh RN (2016).
An ongoing role for Wnt signaling in differentiating melanocytes in vivo. Pigment Cell Melanoma Research doi: 10.1111/pcmr.12568.
  Nichols J, Dowd J, Watson S, Parthasarathy R, Brooks E, Subramanian A, Nachtrab G, Poss KD, Schilling TF, and Kimmel CB. (2016). Repetitive element silencing buffers a mef2ca dependent ligament versus bone fate decision in the zebrafish craniofacial skeleton. Development 143, 4430-4440.
  Schilling TF, Sosnik J and Q Nie (2016). Visualizing retinoic acid morphogen gradients.
In HW Detrich III, M Westerfield and LI Zon (Eds.). The Zebrafish: Cellular and Developmental Biology, Part A Cellular Biology (pp. 139-163). Methods in Cell Biol Vol 133. Elsevier Inc., Academic Press.
  Kawauchi S, Santos R, Muto A, Lopez-Burks M, Schilling TF, Lander AD and Calof AL (2016). Using vertebrate animal models to understand the etiology of developmental defects in Cornelia de Lange Syndrome. Am J Hum Genet C Semin Med Genet doi:10.1002/ajmg.c.31484.
  Sosnik J, Zheng L, Rackauckas CV, Gratton E, Nie Q, and Schilling TF. (2016). Noise modulation of retinoic acid signaling controls sharpening of segmental gene expression boundaries in the zebrafish hindbrain. eLife 5:e14034. doi:10.7554/eLife.14034.
  Le Pabic P, Cooper J and Schilling TF (2016). Developmental basis of phenotypic integration in two Lake Malawi cichlids. EvoDevo 21 7:3 doi: 10.1186/s13227-016-0040-z.
  Subramanian A, and Schilling TF (2015). Tendon development and musculoskeletal assembly: emerging roles for the extracellular matrix. Development 142, 4191-4204. doi: 10.1242/dev.114777.
  Cruz IA, Kappedal R, Mackenzie S, Hoffman TL, Schilling TF and Raible DW (2015). Robust regeneration of adult zebrafish lateral line hair cells reflects continued precursor pool maintenance. Developmental Biology 402, 229-238. doi: 10.1016/j.ydbio.2015.03.019.
  Boer, EF, Howell, ED, Schilling, TF, Jette, CA and Stewart RA (2015). Fascin1-dependent filopodia are required for directional migration of a subset of neural crest cells. PLoS Genetics 11(1):e1004946. doi: 10.1371/journal.pgen.1004946.
  Le Pabic P, Ng C and Schilling TF (2014). Fat-Dachsous signaling coordinates cartilage differentiation and polarity during craniofacial development. PLoS Genetics 10(10):e1004726. doi: 10.1371/journal.pgen.1004726.
  Muto A, Ikeda S, Lopez-Burks ME, Kikuchi Y, Calof AL, Lander AD and Schilling TF (2014). Nipbl and Mediator cooperatively regulate gene expression to control limb development. PLoS Genetics 10(9):e1004671. doi: 10.1371/journal.pgen.1004671.
  Alexander C, Piloto S, Le Pabic P and Schilling TF (2014). Wnt signaling interacts with Bmp and Edn1 to regulate dorsal-ventral patterning and growth of the craniofacial skeleton. PLoS Genetics 10(7):e1004479. doi: 10.1371/journal.pgen.1004479
  Subramanian A and Schilling TF (2014). Thrombospondin-4 controls matrix assembly during development and repair of myotendinous junctions. Elife (Cambridge) doi: 10.7554/eLife.02372.
  Tuttle A, Hoffman T and Schilling TF (2014). rabconnectin-3a regulates vesicle endocytosis and canonical Wnt signaling in zebrafish neural crest migration. PLoS Biology 12(5): e1001852. doi:10.1371/journal.pbio.1001852.
  Schilling TF and Le Pabic P (2014). Neural crest cells in craniofacial skeletal development. In Neural Crest Cell Differentiation and Disease (ed. P Trainor) (pp. 127-151). Elsevier.
  Clemens DM, Nemeth-Cahalan KL, Trinh L, Zhang T, Schilling TF and Hall JE (2013). In vivo analysis of Aquaporin O function in zebrafish: permeability regulation is required for lens transparency. Investigative Ophthalmology and Visual Science doi:pii: iovs.13-12337v1. 10.1167/iovs.13-12337.
  Wu B, Piloto S, Zeng W, Hoverter N, Schilling TF and Waterman M (2013). Ring finger protein 14 is a new regulator of Tcf/B-catenin mediated transcription and colon cancer cell survival. EMBO Reports 14, 347-355. doi: 10.1038/embor.2013.19.
  Schilling TF, Nie Q and Lander AD (2012). Dynamics and precision in retinoic acid morphogen gradients. Current Opinion in Genetics and Development 22, 562-569. doi: 10.1016/j.gde.2012.11.012.
  Nguyen-Chi M, Bryson-Richardson R, Sonntag C, Hall T, Gibson A, Stzal T, Schilling TF and Currie PD (2012). Morphogenesis and cell fate determination within the adaxial cell equivalence group of the zebrafish myotome. PLoS Genetics 8(10):e1003014. doi: 10.1371/journal.pgen.1003014.
  Zhang L*, Radtke K*, Zheng L, Cai AQ, Schilling TF# and Nie Q# (2012). Noise drives sharpening of gene expression boundaries in the zebrafish hindbrain. Molecular Systems Biology 8:613 doi: 10.1038/msb.2012.45. (*co-first authors; # - co-senior authors)
  Cai AQ*, Radtke K*, Linville A, Lander AD, Nie Q# and Schilling TF# (2012). Cellular retinoic acid binding proteins (Crabps) are essential for hindbrain patterning and signal robustness in zebrafish. Development 139, 2150-2155 (*co-first authors - #co-last authors).
  de Pater E, Ciampricotti M, Priller F, Veerkamp J, Strate I, Smith K, Karina A, Schilling TF, Herzog W, Abdelilah-Seyfried S, Hammerschmidt M and Bakkers J (2012). Bmp signaling exerts opposite effects on cardiac differentiation. Circulation Research [Epub ahead of print] PMID 22247485.
  Muto A, Calof A, Lander AD and Schilling TF (2011). Multifactorial origins of heart and gut defects in nipbl-deficient zebrafish: a model of Cornelia de Lange syndrome. PLoS Biol 9(10): e1001181.
  Zuniga E, Rippen M, Alexander C, Schilling TF and Crump JG (2011). Gremlin 2 regulates distinct roles of BMP and Endothelin 1 signaling in dorsoventral patterning of the facial skeleton. Development 138, 5147-5156.
  Alexander C, Zuniga E, Blitz IL, Wada N, Le Pabic P, Javidan Y, Zhang T, Cho KW, Crump JG and Schilling TF (2011) Combinatorial roles for BMPs and Endothelin 1 in patterning the dorsal-ventral axis of the craniofacial skeleton. Development 138, 5135-5146.
  Clouthier D, Garcia E and Schilling TF (2010). Regulation of facial morphogenesis by endothelin signaling: insights from mice and fish. American Journal of Medical Genetics Part A 152A, 2962-2972 (doi:10.1002/ajmg.a.33568).
  Froger A, Clemens D, Kalman K, Nemeth-Cahalan KL, Schilling TF and Hall JE (2010). Two distinct Aquaporin-0s are required for development and transparency of the zebrafish lens. Investigative Ophthalmology and Visual Science 51, 6582-6592.
  Piloto S and Schilling TF (2010). Ovo1 links Wnt signaling with N-cadherin localization during neural crest migration. Development 137, 1981-1990.
  Schilling TF, LePabic P and Hoffman T (2010). Using transgenic zebrafish (Danio rerio) to study development of the craniofacial skeleton. Journal of Applied Ichthyology 26, 187-191.
  Schilling TF and LePabic P (2009). Fishing for the signals that pattern the face. Journal of Biology 8, 101.
  Javidan Y, Alexander C and Schilling TF (2009). Development of cartilage and bone. In "Essential Zebrafish Methods: Cell and Developmental Biology (Reliable Lab Solutions)" (eds. W Dietrich, M Westerfield and L Zon) Academic Press, London. pp. 429-451.
  Linville A, Radtke K, Waxman J, Yelon D and Schilling TF (2009). Combinatorial roles for zebrafish retinoic acid receptors in the hindbrain, limbs and pharyngeal arches. Developmental Biology 325, 60-70.
  Currie P, Schilling TF and Ingham PW (2008). Small scale marker-based screening for mutations in zebrafish development. Methods in Molecular Biology 461, 493-512.
  Nair S and Schilling TF (2008). Chemokine signaling controls endodermal migration during zebrafish gastrulation. Science 322, 89-92.
  White RJ and Schilling TF (2008). How degrading: Cyp26s in hindbrain development. Developmental Dynamics 237, 2775-2790.
  Schilling TF (2008). Anterior-posterior patterning and segmentation of the vertebrate head. J Integrative and Comparative Biology doi:10.1093/icb/icn081.
  Kuratani S and Schilling TF (2008). Head segments in vertebrates. J Integrative and Comparative Biology doi:10.1093/icb/icn036.
  White RJ, Nie Q, Lander AD and Schilling TF (2007). Complex regulation of cyp26a1 creates a robust retinoic acid gradient in the zebrafish embryo. PLoS Biol 5(11): e304:doi:10.13711/journal.pbio.0050304.
  Bryson-Richardson RJ, Berger S, Schilling TF, Hall TE, Cole NJ, Gibson AJ, Sharpe J and Currie PD (2007). FishNet: an online database of zebrafish anatomy. BMC Biology 5:34 doi:10.1186/1741-7007-5-34.
  Hoffman TF, Javier A, Campeau S, Knight RD and Schilling TF (2007). AP2 transcription factors in zebrafish development and vertebrate evolution. J Exp Zool: Mol Dev Evol 308, 679-691.
  Luo T, Xu Y, Hoffman T, Zhang T, Schilling TF and Sargent TD (2007). Inca: a novel p21-activated kinase-associated protein required for cranial neural crest development. Development, 134, 1279-1289.
  Schilling TF (2007). Genetic regulation of neural and non-neural co-evolution. In "Evolution of Nervous Systems" (ed. J. Kaas) Elsevier.
  Nair S, Wei J, Cornell R and Schilling TF (2007). Requirements for endothelin receptors and endothelin-1 signals from facial ectoderm in patterning skeletogenic neural crest in zebrafish. Development, 134, 335-345.
  Webb J and Schilling TF (2006). Zebrafish in comparative context: a symposium. Journal of Integrative and Comparative Biology 46, 569-576.
  Stafford D, White R, Kinkel MD, Linville A, Schilling TF and Prince V (2006). Retinoids signal directly to zebrafish endoderm to specify insulin-expressing beta-cells. Development 133, 949-956.
  Knight RD and Schilling TF (2006). Skeletogenic lineages of the cranial neural crest. In "Neural Crest Induction and Differentiation" (ed. J.-P. Saint-Jeannet) Landes Bioscience; pp. 120-133.
  Wada N., Javidan Y, Nelson S, Carney TJ, Kelsh RN, and Schilling TF (2005). Hedgehog signaling is
required for cranial neural crest morphogenesis and chondrogenesis at the midline in the zebrafish skull.
Development 132, 3977-3988.
  Holzschuh J, Wada N, Wada C, Schaffer A, Javidan Y, Tallafuss A, Bally-Cuif L, and Schilling TF (2005). Requirements for endoderm and BMP signaling in sensory neurogenesis in zebrafish. Development 132, 3731-3742.
  Plaster N, Sonntag C, Schilling TF and Hammerschmidt M (2007) REREa/Atrophin-2 interacts with histone deacetylase and Fgf8 signaling to regulate multiple processes of zebrafish development. Dev Dyn, 236, 1891-1904.
  Norton WH, Mangoli M, Lele Z, Pogoda H-M, Diamond B, Mercurio S, Russell C, Teraoka H, Stickney HL, Rauch G-J, Heisenberg C-P, Houart C, Schilling TF, Frohnhoefer H-G, Rastegar S, Neumann CJ, Gardiner RM, Strahle U, Geisler R, Rees M, Talbot WS and Wilson SW (2005). Monorail/Foxa2 regulates floorplate differentiation and specification of oligodendrocytes, serotonergic raphe neurons and cranial motoneurones. Development 132, 645-658.
  Knight RD, Javidan Y, Zhang T, Nelson S and Schilling TF (2005). AP2-dependent signals from the ectoderm regulate craniofacial development in the zebrafish embryo. Development 132, 3127-3138.
  Javidan Y and Schilling TF (2004). Development of cartilage and bone. Methods Cell Biol. 76, 415-436.
  Clouthier DE and Schilling TF (2004). Understanding Endothelin-1 function during craniofacial development in the mouse and zebrafish. Birth Defects Res. 72, 190-199.
  Linville A, Gumusaneli E, Chandraratna RA and Schilling TF (2004). Independent roles for retinoic acid in segmentation and neuronal differentiation in the zebrafish hindbrain. Dev. Biol. 270, 186-199.
  Knight RD, Javidan Y, Nelson S, Zhang T and Schilling TF (2004). Skeletal and pigment cell defects in the lockjaw mutant reveal multiple roles for zebrafish tfap2a in neural crest development. Dev. Dyn. 229, 87-98.
  Piotrowski T, Ahn D-G, Schilling TF, Nair S, Ruvinsky I, Geisler R, Rauch G-J, Haffter P, Zon LI, Zhou Y, Foott H, Dawid IB and Ho RK (2003). The zebrafish van gogh mutation disrupts tbx1 which is involved in the DiGeorge deletion syndrome in humans. Development 130, 5043-5052.
  Schilling TF (2003). Evolution and development: making jaws. Heredity 90, 3-5.
  Knight RD, Nair S, Nelson S, Javidan Y, Afshar A, Geisler R, Rauch G-J and Schilling TF (2003). The lockjaw mutation reveals requirements for zebrafish tfap2a in early development of the neural crest. Development 130, 5755-5768.
  Yelick PC and Schilling TF (2002). Molecular dissection of craniofacial development using zebrafish. Crit. Rev. Oral Biol. Med. 13, 308-322.
  David N, Saint-Etienne L, Schilling TF and Rosa F (2002). Critical requirement for endoderm and FGF in ventral head skeleton induction. Development 129, 4457-4468.
  Schilling TF (2002). The morphology of larval and adult zebrafish. In Zebrafish: A Practical Approach. (eds. S. Schulte-Merker and C. Nusslein-Volhard) Oxford Univ. Press, Oxford.
  Schilling TF and Knight RD (2001). Origins of anteroposterior patterning and Hox gene regulation during chordate evolution. Proc. R. Soc. Lond. B 268, 1-15
  Begemann G, Schilling TF, Rauch JR, Geisler R and Ingham PW (2001) The zebrafish neckless mutation reveals a requirement for raldh2 in mesodermal signals that pattern the hindbrain. Development 128, 3081-3094.
  Schilling TF, Prince V, and Ingham PW (2001). Plasticity in zebrafish hox expression in the hindbrain and cranial neural crest. Developmental Biology 231, 201-216.
  Miller CT, Schilling TF, Lee K, Parker J and Kimmel CB (2000). sucker encodes a zebrafish Endothelin-1 required for ventral pharyngeal arch development. Development 127, 3815-3828.
  Veitch E, Begbie J, Schilling TF, Smith MM and Graham A (2000). Pharyngeal arch patterning in the absence of neural crest. Current Biology 9, 1481-1484.
  Brown L, Rodaway A, Schilling TF, Jowett T, Ingham PW, Patient R and Sharrocks AD (2000). Insights into vasculogenesis revealed by the expression of the ETS-domain transcription factor Fli-1 in wild-type and mutant zebrafish embryos. Mechanisms of Development 90, 237-252.
  Schilling TF, Concordet J-P and Ingham PW (1999). Regulation of left-right asymmetries in the zebrafish by Shh and BMP4. Developmental Biology 210, 277-287.
  Brown L, Amores A, Schilling TF, Jowett T, Baert JL, de Launoit Y and Sharrocks AD (1998). Molecular characterization of the zebrafish PEA3 ETS-domain transcription factor. Oncogene 17, 93-104.
  Schilling TF and Kimmel CB (1997). Musculoskeletal patterning in the pharyngeal segments of the zebrafish embryo. Development 124, 2945-2960.
  Schilling TF (1997). Genetic analysis of craniofacial development in the vertebrate embryo. Bioessays 19, 459-468.
  Piotrowski T, Schilling TF, Brand M, Jiang Y-J, Heisenberg C-P, Beuchle D, Grandel H, van Eeden FJM, Furutani-Seiki M, Granato M, Haffter P, Hammerschmidt M, Kane DA, Kelsh RN, Mullins MC, Odenthal J, Warga RM and Nusslein-Volhard C (1996). Jaw and branchial arch mutants in zebrafish II: anterior arches and cartilage differentiation. Development 123, 345-356.
  Schilling TF, Piotrowski T, Grandel H, Brand M, Heisenberg C-P, Jiang Y-J, Beuchle D, Hammerschmidt M, Kane DA, Mullins MC, van Eeden FJM, Kelsh RN, Furutani-Seiki M, Granato M, Haffter P, Odenthal J, Warga RM, Trowe T and Nusslein-Volhard C (1996). Jaw and branchial arch mutants in zebrafish I: branchial arches. Development 123, 329-344.
  Schilling TF, Walker C and Kimmel CB (1996). The chinless mutation and cell interactions in zebrafish jaw development. Development 122, 1417-1426.
  Kimmel CB, Ballard WW, Kimmel SR, Ullmann B and Schilling TF (1995). Stages of embryonic development in the zebrafish. Developmental Dynamics 203, 253-310.
  Schilling TF and Kimmel CB (1994). Segment and cell type lineage restrictions during pharyngeal development in the zebrafish embryo. Development 120, 483-494.
  Schilling TF (1993). Cell lineage and mutational analyses of cranial neural crest development in the zebrafish embryo. Ph.D. Thesis. University of Oregon Press, Eugene.
  Thisse C, Thisse B, Schilling TF, and Postlethwaite J (1993). Structure of the zebrafish snail 1 gene and its expression in wild-type, spadetail and notail mutant embryos. Development 119, 1203-1215.
  Kimmel CB, Molven A, Ellis TJ, Hatta K, Ho RK, Kane DA, Schilling TF, Walker C and Warga RM (1993). Developmental and Mutational Analysis of Body Pattern Formation: The Zebrafish Model. In Physiological and Biochemical Aspects of Fish Development. (eds. Walther BT and Fyhn HJ). Grafisk Hus, Bergen, Norway.
  Kimmel CB, Schilling TF and Hatta K (1991). Patterning of body segments of the zebrafish embryo. Current Topics in Developmental Biology 25, 77-110.
  Hatta K, Schilling TF, BreMiller RA, and Kimmel CB (1990). Specification of jaw muscle identity in zebrafish: correlation with engrailed-homeoprotein expression. Science 250, 802-805.
  Kimmel CB, Warga RM and Schilling TF (1990). Origin and organization of the zebrafish fate map. Development 108, 581-594.
Grants “Regulation of morphogenesis and extracellular matrix assembly at the myotendinous junction” Principal Investigator: Thomas Schilling Agency: NIH NIAMS Type: R01 AR67797 Period: 2016 - 2021
“The role of the Fat pathway in cartilage pattern and polarity in zebrafish” Principal Investigator: Thomas Schilling Agency: NIH NIDCR Type: R01 DE13828 Period: 2012 – 2017
“Defining an integrated signaling network that patterns the craniofacial skeleton” Co-PIs: David Clouthier and Thomas Schilling Agency: NIH NICHD Type: R01 HD073182 Period: 2014 – 2019
“Stochastic dynamics and noise control in patterning systems” Co-PIs: Qing Nie and Thomas Schilling Agency: NIH NIGMS Type: R01 GM107264 Period: 2014 – 2018
Society for Developmental Biology
British Society for Developmental Biology
American Association of Anatomists
Graduate Programs Developmental Biology and Genetics

Cellular and Molecular Biosciences

Interdepartmental Neuroscience Program

Mathematical and Computational Biology

Research Centers Developmental Biology Center
Cancer Center
Center for Complex Biological Systems
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Last updated 02/18/2017